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Oxford University Press is a department of the University of Oxford. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide. Sign In or Create an Account. Sign In. Advanced Search. Search Menu. Article Navigation. Close mobile search navigation Article Navigation. Volume Article Contents Abstract. Formation of ent -Kaurene.
Formation of C 20 -Gibberellins. Gibberellin Biosynthesis from GA Regulation of GA Metabolism. Concluding Remarks. Peter Hedden Peter Hedden. Corresponding author: E-mail, peter. Oxford Academic. Select Format Select format.
Permissions Icon Permissions. Abstract Gibberellins are produced by all vascular plants and several fungal and bacterial species that associate with plants as pathogens or symbionts. Fig 1. Open in new tab Download slide. Fig 2. Fig 3. Google Scholar Crossref. Search ADS. Biosynthetic origin of gibberellin A 3 and gibberellin A 7 in cell-free preparations from seeds of Marah macrocarpus and Malus domestica.
Identification and localization of gibberellins in maturing seeds of the cucurbit Sechium edule , and a comparison between this cucurbit and the legume Phaseolus coccineus.
Function and transcript analysis of gibberellin-biosynthetic enzymes in wheat. The biosynthesis of kaurenolide diterpenoids by Gibberella fujikuro I. Fungal products. Characterization of the fungal gibberellin desaturase as a 2-oxoglutarate-dependent dioxygenase and its utilization for enhancing plant growth.
Lipid trafficking at endoplasmic reticulum-chloroplast membrane contact sites. Isolation and characterization of the gibberellin biosynthetic gene cluster in Sphaceloma manihoticola.
Proteomic analysis of the proplastid envelope membrane provides novel insights into small molecule and protein transport across proplastid membranes. Stereochemistry of the metabolic steps from kaurenoic acids to kaurenolides and gibberellins.
Small Grain and Dwarf 2, encoding an HD-Zip II family transcription factor, regulates plant development by modulating gibberellin biosynthesis in rice. OsWOX3A is involved in negative feedback regulation of the gibberellic acid biosynthetic pathway in rice Oryza sativa. Gibberellic acid—a new metabolite from the culture filtrates of Gibberella fujikuroi. A molecular framework for light and gibberellin control of cell elongation. Multiple genes recruited from hormone pathways partition maize diterpenoid defences.
To grow or not to grow: what can we learn on ethylene-gibberellin cross-talk by in silico gene expression analysis? Transport of isoprenoid intermediates across chloroplast envelope membranes. Transcriptional regulation of gibberellin metabolism genes by auxin signaling in arabidopsis. A tandem array of ent -kaurene synthases in maize with roles in gibberellin and more specialized metabolism.
Partial synthesis of ent hydroxyoxonorgibberella-1 10 ,diene-7,dioic acid, a catabolite of gibberellin A 29 , and of related-compounds. Gibberellin metabolism in cell-free-extracts from spinach leaves in relation to photoperiod. Evolutionary analysis of three gibberellin oxidase genes in rice, Arabidopsis, and soybean. Identification and functional analysis of bifunctional ent -kaurene synthase from the moss Physcomitrella patens.
Kaurenolide biosynthesis in a cell-free system from Cucurbita maxima seeds. The CYP88A cytochrome P, ent -kaurenoic acid oxidase, catalyzes three steps of the gibberellin biosynthesis pathway. Arabidopsis ent -kaurene oxidase catalyzes three steps of gibberellin biosynthesis. A plastid envelope location of Arabidopsis ent -kaurene oxidase links the plastid and endoplasmic reticulum steps of the gibberellin biosynthesis pathway.
The involvement of gibberellin oxidase genes in phytochrome-regulated petiole elongation of Arabidopsis. Potential sites of bioactive gibberellin production during reproductive growth in Arabidopsis. Divergence and adaptive evolution of the gibberellin oxidase genes in plants.
A rice semi-dwarf gene, Tan-Ginbozu D35 , encodes the gibberellin biosynthesis enzyme, ent -kaurene oxidase. KNOX action in Arabidopsis is mediated by coordinate regulation of cytokinin and gibberellin activities. The biosynthesis of all major pea gibberellins in a cell-free system from Pisum sativum. The loss of carbon in C 19 -gibberellin biosynthesis in a cell-free system from Pisum sativum L. Where do gibberellin biosynthesis and gibberellin signaling occur in rice plants?
Contribution of the mevalonate and methylerythritol phosphate pathways to the biosynthesis of gibberellins in Arabidopsis. Insight on genes affecting tuber development in potato upon potato spindle tuber viroid PSTVd infection. Evolution and diversity of the 2-oxoglutarate-dependent dioxygenase superfamily in plants. Identification and functional characterization of monofunctional ent -copalyl diphosphate and ent-kaurene synthases in white spruce reveal different patterns for diterpene synthase evolution for primary and secondary metabolism in gymnosperms.
What would be the observable consequences if phospholipid bilayer diffusion of drugs into cells is negligible?
Selective deactivation of gibberellins below the shoot apex is critical to flowering but not to stem elongation of Lolium.
Long-day induction of flowering in Lolium temulentum involves sequential increases in specific gibberellins at the shoot apex. Long-distance transport of phytohormones through the plant vascular system. Touch-induced changes in Arabidopsis morphology dependent on gibberellin breakdown. Ovary-derived precursor gibberellin A 9 is essential for female flower development in cucumber. Functional characterization of gibberellin oxidases from cucumber, Cucumis sativus L.
Expression cloning of a gibberellin oxidase, a multifunctional enzyme involved in gibberellin biosynthesis. Separation and characterization of three 2-oxoglutarate-dependent dioxygenases from Cucurbita maxima L. Regulation of gibberellin oxidase1 expression in spinach by photoperiod. Conserved bases for the initial cyclase in gibberellin biosynthesis: from bacteria to plants. Gibberellin 2-oxidation and the SLN gene of Pisum sativum.
Comprehensive expression analysis of Arabidopsis GA2-oxidase genes and their functional insights. Gibberellin signaling is required for far-red light-induced shoot elongation in Pinus tabuliformis seedlings. Interaction between the eui gene and thermo-sensitive genic male sterility in rice. Structure, function and inhibition of ent-kaurene synthase from Bradyrhizobium japonicum. An ent -kaurene-derived diterpenoid virulence factor from Xanthomonas oryzae pv.
The occurrence of gibberellin A 1 in higher plants—isolation from the seed of runner bean Phaseolus multiflorus. Distribution of gibberellin biosynthetic genes and gibberellin production in the Gibberella fujikuroi species complex. Gibberellins promote nodule organogenesis but inhibit the infection stages of nodulation. Transorganellar complementation redefines the biochemical continuity of endoplasmic reticulum and chloroplasts. Gibberellin oxidase activities in Bradyrhizobium japonicum bacteroids.
Gibberellin biosynthesis in bacteria: separate ent -copalyl diphosphate and ent -kaurene synthases in Bradyrhizobium japonicum. Functional characterization of two class II diterpene synthases indicates additional specialized diterpenoid pathways in maize Zea mays. Mechanism of prolyl hydroxylase reaction.
Investigating the phylogenetic range of gibberellin biosynthesis in bacteria. An operon for production of bioactive gibberellin A 4 phytohormone with wide distribution in the bacterial rice leaf streak pathogen Xanthomonas oryzae pv. DELLAs control plant immune responses by modulating the balance and salicylic acid signaling.
Gibberellin hormone signal perception: down-regulating DELLA repressors of plant growth and development. Characterization of CYP as a gibberellin 3-oxidase indicates that certain rhizobia can produce bioactive gibberellin A 4.
Labeling studies clarify the committed step in bacterial gibberellin biosynthesis. Elucidation of gibberellin biosynthesis in bacteria reveals convergent evolution. A modified micro-drop bioassay using dwarf rice for detection of femtomol quantities of gibberellins.
Isolation and expression profiles of gibberellin metabolism genes in developing male and female cones of Pinus tabuliformis. Heterologous expression and transcript analysis of gibberellin biosynthetic genes of grasses reveals novel functionality in the GA3ox family.
Molecular characterization of Rht-1 dwarfing genes in hexaploid wheat. Uncovering the complex metabolic network underlying diterpenoid phytoalexin biosynthesis in rice and other cereal crop plants. Analysis of the developmental roles of the Arabidopsis gibberellin oxidases demonstrates that GA20OX1, -2, and -3 are the dominant paralogs.
Synergistic substrate inhibition of ent -copalyl diphosphate synthase: a potential feed-forward inhibition mechanism limiting gibberellin metabolism.
Gibberellin concentration and transport in genetic lines of pea—effects of grafting. Chemical regulators of gibberellin status and their application in plant production. The gibberellin precursor GA 12 acts as a long-distance growth signal in Arabidopsis. Gibberellin 3-oxidase gene expression patterns influence gibberellin biosynthesis, growth, and development in pea.
The gibberellin biosynthetic genes AtGA20ox1 and AtGA20ox2 act, partially redundantly, to promote growth and development throughout the Arabidopsis life cycle. The makings of a gradient: spatiotemporal distribution of gibberellins in plant development. In vivo gibberellin gradients visualized in rapidly elongating tissues. The P gene of Gibberella fujikuro i encodes a multifunctional enzyme in gibberellin biosynthesis.
Interactions between brassinosteroids and gibberellins: synthesis or signaling? Expression of a gibberellin 2-oxidase gene around the shoot apex is related to phase transition in rice. An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Hydrolysis and reconjugation of gibberellin A 20 glucosyl ester by seedlings of Zea mays L.
The role of a class III gibberellin 2-oxidase in tomato internode elongation. Molecular evolution of the substrate specificity of ent -kaurene synthases to adapt to gibberellin biosynthesis in land plants. Developmental regulation of the gibberellin biosynthetic gene GA1 in Arabidopsis thaliana.
Gibberellin A 1 biosynthesis in Pisum sativum L. Hormonal regulation of temperature-induced growth in Arabidopsis. Comprehensive analysis of cucumber gibberellin oxidase family genes and functional characterization of CsGA20ox1 in root development in Arabidopsis. The Arabidopsis GA1 locus encodes the cyclase ent -kaurene synthetase A of gibberellin biosynthesis.
Google Scholar PubMed. A common allosteric mechanism regulates homeostatic inactivation of auxin and gibberellin. Antheridiogen determines sex in ferns via a spatiotemporally split gibberellin synthesis pathway. DELLA proteins: master regulators of gibberellin-responsive growth and development.
Molecular cloning and functional expression of gibberellin 2-oxidases, multifunctional enzymes involved in gibberellin deactivation. The P gene of Gibberella fujikuroi encodes ent -kaurene oxidase in the gibberellin biosynthesis pathway. Characterization of the final two genes of the gibberellin biosynthesis gene cluster of Gibberella fujikuroi des and P encode GA 4 desaturase and the hydroxylase, respectively.
The gibberellin oxidase of Gibberella fujikuroi is a multifunctional monooxygenase. Geranyl diphosphate synthase is required for biosynthesis of gibberellins. CYPA8: a rice ent-kaurene oxidase paralog diverted to more specialized diterpenoid metabolism. Gibberellin 3-oxidases in developing embryos of the southern wild cucumber, Marah macrocarpus. Probing the mechanism of loss of carbon in gibberellin biosynthesis. Stamens and gibberellin in the regulation of corolla pigmentation and growth in Petunia hybrida.
Deciphering the cryptic genome: genome-wide analyses of the rice pathogen Fusarium fujikuroi reveal complex regulation of secondary metabolism and novel metabolites. Functional characterization of wheat copalyl diphosphate synthases sheds light on the early evolution of labdane-related diterpenoid metabolism in the cereals.
GAMT2 encodes a methyltransferase of gibberellic acid that is involved in seed maturation and germination in Arabidopsis. The dioxygenase GIM2 functions in seed germination by altering gibberellin production in Arabidopsis. Functional characterization of the rice kaurene synthase-like gene family. Gibberellin acts positively then negatively to control onset of flower formation in Arabidopsis. Molecular cloning and characterization of a cDNA encoding the gibberellin biosynthetic enzyme ent -kaurene synthase B from pumpkin Cucurbita maxima L.
The GA2 locus of Arabidopsis thaliana encodes ent -kaurene synthase of gibberellin biosynthesis. The methyl ester of a new gibberellin, GA 73 —the principal antheridiogen in Lygodium japonicum. Mutations on ent -kaurene oxidase 1 encoding gene attenuate its enzyme activity of catalyzing the reaction from ent -kaurene to ent -kaurenoic acid and lead to delayed germination in rice.
The gibberellin oxidase that specifically converts gibberellin A 9 to A 20 in Tripterygium wilfordii is a 2-oxoglutarate-dependent dioxygenase. A recruiting protein of geranylgeranyl diphosphate synthase controls metabolic flux toward chlorophyll biosynthesis in rice. Investigating the conservation pattern of a putative second terpene synthase divalent metal binding motif in plants.
Functional characterization of wheat ent -kaurene -like synthases indicates continuing evolution of labdane-related diterpenoid metabolism in the cereals. To gibberellins and beyond! Issue Section:. Download all slides. View Metrics. Email alerts Article activity alert. Advance article alerts.
Figure 1. Simplified GA metabolic pathway in flowering plants. All enzymes mapped to the GA metabolic pathway, except GA13ox from Tripterygium wilfordii , were verified with enzymatic assays and the chemical profiling of transgenic plants. Ps ER membrane-associated proteins in this pathway are highlighted in blue, while 2-oxoglutarate-dependent dioxygenases 2-ODDs, cytosolic proteins are highlighted in purple.
All dashed lines indicate enzymatic a step that has not yet been characterized at the genetic level in any flowering plant. The Arabidopsis GA-deficient mutants ga1 to ga5 used for map-based cloning are shown in parentheses next to the corresponding genes. The GA metabolic pathway at the molecular level has been extensively investigated using both forward and reverse genetics strategies.
Both strategies used on GA metabolism started with the dwarfism phenotype caused by bioactive GA deficiency e. A forward genetics strategy, in which a mutant population is scored by the dwarf phenotype change followed by map-based cloning of the causal genes, has been applied to GA biosynthesis gene identification in Arabidopsis, from GGPP to GA 4 , for a long time.
Thanks to the work of phytochemists, naturally occurring GAs have been identified thus far Hedden, However, no novel GAs have been reported in plants for a long time. Recently, Liu et al. Furthermore, DHGA 12 or GA 4 could only partially rescue the phenotype of cotyledon greening and hypocotyl elongation in gas2 mutants, indicating that there were other factors involved in these processes.
Comprehensive metabolite analysis of GAS2 transgenic lines will provide valuable evidence to elucidate the GAS2 functions in Arabidopsis. Moreover, the resulting transgenic plants showed a substantial growth increase, which reflected the increase in the total bioactivity of GA in planta Bhattacharya et al. However, the functional homologs of fungal GA 4 -DES have not been characterized from plants to date.
Previous studies have shown that not only GA biosynthesis but also GA catabolism deactivation play important roles in GA bioactivity homeostasis in planta. It is plausible to speculate that other members of the CYP family might be involved in GA deactivation in plants. Figure 2. Characterization of GA oxidases encoded by P genes in flowering plants. P proteins with GA oxidase activity are marked with red circles. Data were extracted from the Tomato Genome Consortium Sato et al.
Data were extracted from Shen et al. Shen et al. Moreover, CYP72A9 was expressed predominantly in developing seeds in Arabidopsis , and cyp72a9 mutants showed a deficiency of GA 1 but an increase of GA 4 in the siliques. CYP72A9 was further proven to be involved in primary seed dormancy He et al. The other three Arabidopsis CYP72A members A7, A13, and A15 also showed GA hydroxylase activity, but overexpression plants did not show a semi-dwarf phenotype, which merits further investigation of the physiological functions of these genes in planta.
As both CYP and CYP72A are widely distributed in flowering plants, it will be interesting to determine the role of each member of both P subfamilies in contributing to GA homeostasis in different plant species Figure 2.
Thus, further investigation is needed to test whether CYP72A is involved in leaf development in soybean. Zhang et al. TwGA13ox was highly expressed in root phloem and could be induced by methyl jasmonate MeJA treatment. It is noteworthy that the more detailed biochemical and physiological function of Tw GA13ox has not yet been determined in Tripterygium wilfordii.
Lange et al. However, it is unlikely that 2-ODD enzymes are the main contributor to OH GA production in flowering plants, as the above-mentioned Ps are responsible for OH GA production in rice and Arabidopsis, which has been shown at the biochemical and genetic levels. Due to the very low level of endogenous GAs, accurate quantitative measurement plays an important role in elucidating the GA metabolic pathway in plants. Currently, triple quadrupole MS mass spectrometry with MRM multi-reaction monitoring is the most frequently used operation mode for the quantitative analysis of target GAs Pan et al.
The purification of a novel GA from plants and the elucidation of its absolute chemical structure is always a time-consuming and laborious procedure. Since GAs play important roles in regulating plant growth and development, the GA genes described here represent a promising toolkit with which synthetic biologists can adjust bioactive GA levels to manipulate plant traits. The constitutive promoters, coupled with additional copies of the GA genes, were applied in all of the above-mentioned cases in Arabidopsis.
All these results also confirmed that GA genes are sensitive to endogenous bioactive GA changes: GA biosynthesis genes are usually upregulated when GA 4 decreases, while GA catabolism genes are downregulated under the same conditions, and vice versa He et al. At the enzyme level, the targeted near-saturated mutagenesis of GA genes, especially those downstream of GA 12 formation multiple copies with different tissue-specificity , using CRISPR-based editing could be applied to fine-tune bioactive GA levels to achieve ideal plant traits Chen K.
After quantitatively characterizing these GA components in planta , other GA-related factors, such as signal transduction and interactions with other phytohormones, etc.
GW and JH conceptualized the content of the manuscript. JH wrote the first draft of the manuscript. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Beck, G.
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